Balto-Slavs and Sarmatians in the Battle of Himera

G25 coordinates for most of the samples from the recent Reitsema et al. paper are available in a text file here. They're also in the G25 datasheets at the usual link here.

A basic distance analysis with the G25 data at Vahaduo shows that the two samples labeled Himera_480BCE_3 are either early Balts or Slavs. I suspect that they're Slavs, because I believe that early Slavs had this type of Baltic-like genetic structure before mixing with their non-Slavic-speaking neighbors. Well, that's my pet theory for now, so take it or leave it.

Distance to: ITA_Sicily_Himera_480BCE_3:I10943
0.03393838 HUN_IA_La_Tene_o:I18226
0.03572886 DEU_MA_Krakauer_Berg:KRA001
0.03618075 RUS_Pskov_VA:VK159
0.03899963 SWE_Gotland_VA:VK463
0.03915018 Baltic_EST_IA:s19_V12_1

Distance to: ITA_Sicily_Himera_480BCE_3:I10949
0.03573636 HUN_IA_La_Tene_o3:I25524
0.03698768 HUN_IA_La_Tene_o:I18226
0.03732752 SWE_Skara_VA:VK397
0.03767022 Baltic_EST_IA:s19_V12_1
0.03772687 DEU_MA_Krakauer_Berg:KRA001

On the other hand, I'm almost certain that the two Himera_480BCE_4 samples are Sarmatians. The good old G25 does it again!

Distance to: ITA_Sicily_Himera_480BCE_4:I10944
0.03100861 KAZ_Segizsay_Sarmatian:SGZ002
0.03548059 MDA_Sarmatian:I11925
0.03619219 RUS_Urals_Sarmatian:MJ56
0.03626538 RUS_Urals_Sarmatian:chy001
0.03904260 RUS_Urals_Sarmatian:MJ41

Distance to: ITA_Sicily_Himera_480BCE_4:I10947
0.02989458 RUS_Urals_Sarmatian:MJ43
0.03052790 RUS_Urals_Sarmatian:chy002
0.03170622 KAZ_Kangju:DA226
0.03288789 TUR_BlackSea_Samsun_Anc_C:I4529
0.03310149 KAZ_Aigyrly_Sarmatian:AIG003

See also...

Slavic-like Medieval Germans

Fatyanovo as part of the wider Corded Ware family (Nordqvist and Heyd 2020)

There's a new archeological paper about the Fatyanovo culture at the Proceedings of the Prehistoric Society [LINK]. It includes this quote on page 18:

In the traditional narrative, the Fatyanovo people – like the CWC populations in general – are regarded as Indo-European, representing the pre-Balto-Slavic (-Germanic) stage (Carpelan & Parpola 2001, 88; Anthony 2007, 380; also Gimbutas 1956, 163; Tretyakov 1966, 109) in the spread of Indo-European languages.

That's correct, but considering the latest ancient DNA research on the Fatyanovo people, the traditional narrative is probably wrong. Fatyanovo males were rich in Y-haplogroup R1a-Z93, which is found at very low frequencies in Balto-Slavic populations (see here). It's actually much more common nowadays in Central and South Asia, where it often reaches frequencies of over 50% in Indo-Iranian speaking groups.

Balts and Slavs are rich in R1a-Z282, which is a sister clade of R1a-Z93 that has been found in Corded Ware and Corded Ware-related samples from west of Fatyanovo sites. That is, in present-day Poland and the Baltic states.

Therefore, the origins of the Balto-Slavs should be sought somewhere west of the Fatyanovo culture, probably in the Corded Ware derived populations from what is now the border zone between Poland, Belarus and Ukraine.

Indeed, in my view the Fatyanovo people are more likely to have spoken Proto-Indo-Iranian rather than anything ancestral to Baltic or Slavic (see here).

Nordqvist and Heyd, The Forgotten Child of the Wider Corded Ware Family: Russian Fatyanovo Culture in Context, Proceedings of the Prehistoric Society, online 12 November 2020, DOI: https://doi.org/10.1017/ppr.2020.9

See also...

The oldest R1a to date

Big deal of 2019: ancient DNA confirms the link between Y-haplogroup N and Uralic expansions

The academic consensus is that Indo-European languages first spread into the Baltic region from the Eastern European steppes along with the Corded Ware culture (CWC) and its people during the Late Neolithic, well before the expansion of Uralic speakers into Fennoscandia and surrounds, probably from somewhere around the Ural Mountains.

On the other hand, the views that the Uralic language family is native to Northern Europe and/or closely associated with the CWC are fringe theories usually espoused by people not familiar with the topic or, unfortunately it has to be said, mentally unstable trolls.

The likely close relationship between the CWC expansion and the early spread of Indo-European languages was discussed in several papers in recent years (for instance, see here). This year, we saw the first ancient DNA paper focusing on the transition from the Bronze Age to the Iron Age in the East Baltic, including the likely first arrival of Uralic speech in what is now Estonia.

Published in Current Biology courtesy of Saag et al., the paper showed that the genetic structure of present-day East Baltic populations largely formed in the Iron Age (see here). It was during this time, the authors revealed, that the region experienced a sudden influx of Y-chromosome haplogroup N, which is today common in many Uralic speaking populations and often referred to as a Proto-Uralic marker. Little wonder then that Saag et al. linked this genetic shift in the East Baltic to the westward migrations of early Uralic speakers.

The table below, based on data from the Saag et al. paper, surely doesn't leave much to the imagination about what happened.

Unfortunately, I have to say that the genome-wide analysis in the paper was less informative than it could have been. The authors focused their attention on rather broad genetic components, and, as a result, missed an interesting fine scale distinction between their Bronze Age and Iron Age samples. The spatial maps below, based on my Global25 data for most of the ancients from Saag et al., show what I mean. The hotter the color the higher the genetic similarity between them and present-day West Eurasian populations.

Note that the Bronze Age (Baltic_EST_BA) samples are most similar to the Baltic-speaking, and thus also Indo-European-speaking, Latvians and Lithuanians, rather than the Uralic-speaking Estonians, even though they're from burial sites in Estonia. On the other hand, the Iron Age (Baltic_EST_IA) samples show strong similarity to a wider range of populations, including Estonians and many other Uralic-speaking groups.

See also...

It was always going to be this way

Fresh off the sledge

More on the association between Uralic expansions and Y-haplogroup N

Fresh off the sledge

As things stand, the closest individual to a Proto-Uralic speaker in the ancient DNA record is arguably 0LS10 from an Iron Age tarand grave in what is now Estonia. I say that because:

- isotopic data suggest that 0LS10 wasn't born where he died, and considering his elevated Siberian ancestry relative to earlier and most contemporaneous Baltic ancients, he was very likely a migrant to the Baltic region from the east

- the tarand grave tradition appears to be specifically a Finnic (west Uralic) phenomenon that probably spread from the Volga-Oka region, which is just west of where most people place the Proto-Uralic homeland

- 0LS10 belongs to Y-chromosome haplogroup N-L1026, a paternal marker that is especially closely associated with Uralic-speaking populations and probably only appeared in the East Baltic region during the transition from the Bronze Age to the Iron Age

You can find more background info about 0LS10 and other relevant samples in Saag et al. 2019 (see here). This is where he sits in my Principal Component Analyses (PCA) focusing on fine scale Northern European genetic diversity. The relevant datasheets are available here and here, respectively.

Note that 0LS10 doesn't cluster strongly with any ancient or modern populations. To investigate this in more detail I ran a series of two-way qpAdm analyses, testing tens of ancient individuals and populations as potential admixture sources. These two models stood out above the rest in terms of their statistical fits, chronology and overall plausibility.

Baltic_EST_IA_0LS10
Baltic_EST_BA 0.826±0.045
RUS_Sintashta_MLBA_o1 0.174±0.045
chisq 12.527
tail prob 0.564048
Full output

Baltic_EST_IA_0LS10
Baltic_EST_BA 0.683±0.102
RUS_Mezhovskaya 0.317±0.102
chisq 13.811
tail prob 0.463864
Full output

Please note that RUS_Sintashta_MLBA_o1 isn't representative of the Sintashta culture population as a whole. It's a group of the most extreme genetic outliers among the Sintashta samples, and they may or may not have been Uralic speakers (see here). Interestingly, the Mezhovskaya culture population is generally associated with the Ugric branch of the Uralic language family.

I was also able to closely replicate these results with the Global25/nMonte method; down to almost one per cent. However, the statistical fits (distances) are poor, probably because the reference populations aren't the real mixture sources. This is in line with the fact that their Y-haplogroups are Q1a, R1a and R1b, rather than any type of N.

Baltic_EST_IA:0LS10
Baltic_EST_BA,83.8
RUS_Sintashta_MLBA_o1,16.2
distance%=4.7955

Baltic_EST_IA:0LS10
Baltic_EST_BA,69.8
RUS_Mezhovskaya,30.2
distance%=3.5783

I do realize that two Bronze Age samples from Bolshoy Oleni Ostrov, Kola Peninsula, belong to N-L1026, but adding them to my mixture models doesn't help. Little wonder, because the Kola Peninsula lies within the Arctic Circle, and I'm pretty sure that 0LS10 and his N-L1026 came from somewhere just north of the mixture cline marked on the map below. Unfortunately, I can't test this directly yet due to the scarcity of ancient samples from this region.

See also...

It was always going to be this way

On the association between Uralic expansions and Y-haplogroup N

Uralic-specific genome-wide ancestry did make a signifcant impact in the East Baltic

Uralic-specific genome-wide ancestry did make a signifcant impact in the East Baltic

I've started analyzing the ancient genotype data from the recent Saag et al. paper on the expansion of Uralic languages and associated spread of Siberian ancestry into the East Baltic region. The paper is freely available here and the data are here.

I really like the paper, but I don't agree with the authors' claim that the appearance of Y-chromosome haplogroup N in what is now Estonia and surrounds during the Iron Age is "not matched by a clear shift in autosomal profiles". In my opinion it certainly is, and, as one would expect, it's a shift towards a genetic profile typical of western Uralic speakers.

I'd say that the easiest way to find this signal is with a Principal Component Analysis (PCA) focusing on fine scale genetic substructures within Northern Europe, like the one below. The relevant datasheet is available here.

Note that the East Baltic Iron Age samples, all from burial sites in what is now Estonia, appear to be peeling away from their Bronze Age predecessors and overlapping strongly with present-day Estonians, who are Uralic speakers. Indeed, the PCA suggests to me that the formation of the greater part of the present-day Estonian gene pool took place in the East Baltic during the transition from the Bronze Age to the Iron Age. That is, when Uralic languages are generally accepted to have arrived in the region from near the Ural Mountains in the east.

I was also able to closely replicate these outcomes with my Global25 data using the method described here. However, in this effort, present-day Estonians are clearly more western than the Estonian Iron Age samples (EST_IA), which might be due to the presence of low level Germanic ancestry in Estonia dating to the medieval period. The relevant datasheet is available here.

Interestingly, the Estonian Bronze Age samples (EST_BA) come from stone-cist graves which are widely hypothesized to have been introduced to the East Baltic from the Nordic Bronze Age civilization. I even recall reading a paper on the topic which claimed that the remains buried in such graves were those of Proto-Germanic-speaking Scandinavian migrants. Well, I haven't had a chance to study these samples in any great detail yet, but considering that in both of the PCA above they're overlapping strongly with Latvian Bronze Age samples (LVA_BA) and sitting far away from the nearest Scandinavians, I'd say they're probably of local stock from way back.

See also...

It was always going to be this way

On the association between Uralic expansions and Y-haplogroup N

Inferring the linguistic affinity of long dead and non-literate peoples: a multidisciplinary approach

It was always going to be this way

The native peoples of the East Baltic - Estonians, Latvians and Lithuanians - are genetically alike and their paternal gene pools are dominated by the same two Y-chromosome haplogroups: R1a and N3a.

Linguistically, however, Estonians are a world apart from Latvians and Lithuanians. That's because the Estonian language belongs to the Uralic language family, which has an obvious North Eurasian character. On the other hand, Latvian and Lithuanian are both classified as Indo-European languages, along with the vast majority of other European languages.

The Uralic and Indo-European language families may or may not descend from the same ancestral tongue, but even if they do, their relationship is very distant.

So how is it that Estonians came to speak a Uralic language? As far back as I can remember, the basic explanation accepted by most people was that Uralic speech arrived in what is now Estonia and neighboring Finland during the Bronze Age with migrants, or perhaps invaders, rich in N3a from somewhere around the Ural Mountains. Conversely, Latvians and Lithuanians were generally assumed to have retained the Indo-European speech of their R1a-rich forefathers from the Pontic-Caspian steppe, who colonized much of Eastern Europe north of the steppe during the Late Neolithic.

Ancient DNA has now uncannily corroborated these theories (for instance, see Mittnik et al. 2018 and, published today, Saag et al. 2019). All it took was a handful of samples from a few relevant sites. I think that's awesome; I love it when sensible, long-standing hypotheses are validated by cutting edge science.

I'll have a lot more to say about the spread of Uralic languages and Uralian genes to the East Baltic when I get my hands on the genotype data from the new Saag et al. paper. I also have a post coming soon about the Nordic Bronze Age. Stay tuned.

Update 10/05/2019: Uralic-specific genome-wide ancestry did make a signifcant impact in the East Baltic

See also...

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

Corded Ware people =/= Proto-Uralics (Tambets et al. 2018)

Inferring the linguistic affinity of long dead and non-literate peoples: a multidisciplinary approach

So who's the most (indigenous) European of us all?

Basically, the first map below reveals the answer. It shows the spread of a European specific cluster from a global-wide ADMIXTURE analysis at K=8 (eight ancestral populations assumed), which I call "North European". Thus, genetically, the most European populations are found around the Baltic Sea, and in particular in the East Baltic region. In my genome collection, samples from Lithuania clearly and consistently score the highest percentages in ADMIXTURE clusters specific to Europe. However, I suspect that if I had Latvians with no known foreign ancestry going back more than four generations, they'd come out the "most European". Hopefully we can test that in the near future.

Below are the fifteen Eurogenes sample sets that scored the highest levels of membership in the North European cluster. The list only includes groups with five or more individuals present in the analysis, so some populations, like Estonians or Danes, weren't included, even though they easily made the cut. The spreadsheet with all the results from this run can be seen here. A table of Fst (genetic) distances between the eight clusters is available here.

Lithuanians 77%
Finns 74%
Belorussians 70%
Swedes 69%
Norwegians 68%
Kargopol Russians 68%
Russians 68%
Poles 68%
Erzya 66%
Ukrainians 66%
Moksha 66%
Orcadians 63%
HapMap Utah Americans (CEU) 63%
Irish 63%
British 62%

So why did I pick the results from K=8, and not some other K, like 2, 10, or 25? Well, it's not possible to evaluate who is more European without a European-specific cluster (ie. modal in Europeans, with a low frequency outside of Europe). Provided that a decent number and range of global and West Eurasian samples are used in the analysis, such clusters begin appearing at around K=5 or K=6, and start breaking up into local clusters from about K=9. I found that runs below K=8 produced European clusters that spilled too generously outside of the borders of Europe. On the other hand, runs above K=8 produced European clusters that weren't representative of enough European groups (ie. too localized). But the European cluster from K=8 was pretty much perfect, and I think that's obvious from the map. In fact, I can hardly believe how well it fits the modern geographic concept of Europe - north of the Mediterranean and west of the Urals. Amazing stuff.

There are two other clusters that show up across Europe in non-trivial amounts - Mediterranean and Caucasus (see maps below). These can also be thought of as native European clusters, since they've been on the continent for thousands of years. However, their peak frequencies are found in West Asia, so they're not particularly useful signals of European-specific ancestry.

So what do these three clusters show exactly? They represent certain allele frequencies in modern populations, and in fact, these can change fairly rapidly due to admixture, selection, and genetic drift. So claiming that such clusters represent pure ancient populations is unlikely to be true in most cases, if ever. However, I don't think there's anything wrong in saying that, when robust enough, they can be thought of as signals of ancestry from relatively distinct ancestral groups.

Indeed, anyone who's read up on the prehistory of Europe, knows that there are three general Neolithic archeological waves to consider when trying to untangle the story of the peopling of Europe. These are Mediterranean Neolithic, Anatolian Neolithic and Forest Neolithic (for example, see here).

Mediterranean Neolithic refers to a series of migrations from West Asia via the Mediterranean and its coasts. The areas most profoundly affected by these movements include the islands of Sardinia and Corsica, and the Southwest European mainland. Anatolian Neolithic describes migrations into Europe from modern day Turkey, mostly into the Balkans, but also as far as Germany and France. At the moment, Forest Neolithic of Northeastern Europe is something of a mystery. However, the general opinion is that it was largely the result of native Mesolithic hunter-gatherers adopting agriculture.

Obviously, it's very difficult to dismiss the correlations between these three broad archeological groups and the European and two European/West Asian clusters produced in my K=8 ADMIXTURE analysis. Is it a coincidence that the Mediterranean cluster today peaks in Sardinia, which has been largely shielded from foreign admixture since the Neolithic, and today forms a very distinct Southern European isolate? Why does the North European cluster show the highest peaks in classic Forest Neolithic territory? And why does the Caucasus cluster radiate in Europe from the southeast, which is where Anatolian farmers had the greatest impact? These can't all be coincidences, and I'm willing to bet that none of them are. I'm convinced that the three clusters from my K=8 run are strong signals from the Neolithic, and the North European cluster also from the Mesolithic.

Eventually, these issues will be settled with ancient DNA data, in a much more comprehensive way than ever possible using modern genomes. We've already seen some preliminary results, mostly from Mesolithic, Neolithic and Bronze Age sites around Europe, so perhaps it's useful to ask whether my ADMIXTURE analysis and commentary here mirror these early findings? I think they do. For instance, here's an interesting conclusion regarding the East Baltic area from a study on ancient Scandinavian mtDNA by Malmström et al.

Through analysis of DNA extracted from ancient Scandinavian human remains, we show that people of the Pitted Ware culture were not the direct ancestors of modern Scandinavians (including the Saami people of northern Scandinavia) but are more closely related to contemporary populations of the eastern Baltic region. Our findings support hypotheses arising from archaeological analyses that propose a Neolithic or post-Neolithic population replacement in Scandinavia [7]. Furthermore, our data are consistent with the view that the eastern Baltic represents a genetic refugia for some of the European hunter-gatherer populations.

I suppose there will be people wondering why I didn't take Sub-Saharan African, East Asian, and South Asian admixtures into account in my analysis. The reason is that I wasn't looking at which group was most West Eurasian, or Caucasoid. Based on everything I've seen to date, in my own work as well as elsewhere, the most West Eurasian group would probably be the French Basques from the HGDP. However, the differences between them, and certain groups from Northeastern Europe, like Northern Poles and Lithuanians, really wouldn't be that great anyway. I might do a write up about that at some point.


Credits...

- Maps by Eurogenes project member FR7

- Additional stats by Eurogenes project member DESEUK1


References...

Helena Malmström et al., Ancient DNA Reveals Lack of Continuity between Neolithic Hunter-Gatherers and Contemporary Scandinavians, Current Biology, 24 September 2009, doi:10.1016/j.cub.2009.09.017

Noreen von Cramon-Taubadel and Ron Pinhasi, Craniometric data support a mosaic model of demic and cultural Neolithic diffusion to outlying regions of Europe, Proc. R. Soc. B published online 23 February 2011, doi: 10.1098/rspb.2010.2678